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E2F4 regulates transcriptional activation in mouse embryonic stem cells independently of the RB family.
E2F transcription factors are central regulators of cell division and cell fate decisions. E2F4 often represents the predominant E2F activity in cells. E2F4 is a transcriptional repressor implicated in cell cycle arrest and whose repressive activity depends on its interaction with members of the RB family. Here we show that E2F4 is important for the proliferation and the survival of mouse embryonic stem cells. In these cells, E2F4 acts in part as a transcriptional activator that promotes the expression of cell cycle genes. This role for E2F4 is independent of the RB family. Furthermore, E2F4 functionally interacts with chromatin regulators associated with gene activation and we observed decreased histone acetylation at the promoters of cell cycle genes and E2F targets upon loss of E2F4 in RB family-mutant cells. Taken together, our findings uncover a non-canonical role for E2F4 that provide insights into the biology of rapidly dividing cells
An ALMA Survey of H₂CO in Protoplanetary Disks
H₂CO is one of the most abundant organic molecules in protoplanetary disks and can serve as a precursor to more complex organic chemistry. We present an Atacama Large Millimeter/submillimeter Array survey of H₂CO toward 15 disks covering a range of stellar spectral types, stellar ages, and dust continuum morphologies. H₂CO is detected toward 13 disks and tentatively detected toward a fourteenth. We find both centrally peaked and centrally depressed emission morphologies, and half of the disks show ring-like structures at or beyond expected CO snowline locations. Together these morphologies suggest that H₂CO in disks is commonly produced through both gas-phase and CO-ice-regulated grain-surface chemistry. We extract disk-averaged and azimuthally-averaged H₂CO excitation temperatures and column densities for four disks with multiple H₂CO line detections. The temperatures are between 20–50 K, with the exception of colder temperatures in the DM Tau disk. These temperatures suggest that H₂CO emission in disks generally emerges from the warm molecular layer, with some contributions from the colder midplane. Applying the same H₂CO excitation temperatures to all disks in the survey, we find that H₂CO column densities span almost three orders of magnitude (~5 × 10¹¹–5 × 10¹⁴ cm⁻²). The column densities appear uncorrelated with disk size and stellar age, but Herbig Ae disks may have less H₂CO compared to T Tauri disks, possibly because of less CO freeze-out. More H₂CO observations toward Herbig Ae disks are needed to confirm this tentative trend, and to better constrain under which disk conditions H₂CO and other oxygen-bearing organics efficiently form during planet formation
Conséquence d'une coupe rase ou d'une éclaircie sur la richesse spécifique et le mode de dissémination des espèces végétales dans des forêts de pin d'Alep du Var (Sud de la France)
Les conséquences de différents types de coupe sur la diversité végétale et le suivi des variations observées dans le temps ont été analysées par l'intermédiaire de la richesse spécifique et d'une approche plus fonctionnelle sur des forêts de pin d'Alep du Centre Var (variations du nombre d'espèces et des modes de dissémination des espèces végétales). Les résultats indiquent que les effets de ces différents types de coupe sont très marqués les trois premières années après la coupe, puis ils s'estompent jusqu'à pratiquement disparaître au bout de dix ans environ
An ALMA Survey of H₂CO in Protoplanetary Disks
H₂CO is one of the most abundant organic molecules in protoplanetary disks and can serve as a precursor to more complex organic chemistry. We present an Atacama Large Millimeter/submillimeter Array survey of H₂CO toward 15 disks covering a range of stellar spectral types, stellar ages, and dust continuum morphologies. H₂CO is detected toward 13 disks and tentatively detected toward a fourteenth. We find both centrally peaked and centrally depressed emission morphologies, and half of the disks show ring-like structures at or beyond expected CO snowline locations. Together these morphologies suggest that H₂CO in disks is commonly produced through both gas-phase and CO-ice-regulated grain-surface chemistry. We extract disk-averaged and azimuthally-averaged H₂CO excitation temperatures and column densities for four disks with multiple H₂CO line detections. The temperatures are between 20–50 K, with the exception of colder temperatures in the DM Tau disk. These temperatures suggest that H₂CO emission in disks generally emerges from the warm molecular layer, with some contributions from the colder midplane. Applying the same H₂CO excitation temperatures to all disks in the survey, we find that H₂CO column densities span almost three orders of magnitude (~5 × 10¹¹–5 × 10¹⁴ cm⁻²). The column densities appear uncorrelated with disk size and stellar age, but Herbig Ae disks may have less H₂CO compared to T Tauri disks, possibly because of less CO freeze-out. More H₂CO observations toward Herbig Ae disks are needed to confirm this tentative trend, and to better constrain under which disk conditions H₂CO and other oxygen-bearing organics efficiently form during planet formation
An unbiased ALMA spectral survey of the LkCa 15 and MWC 480 protoplanetary disks
Funding: R.A.L. gratefully acknowledges funding from ALMA Student Observing Support and a Jansky Fellowship. K.I.O. acknowledges funding from the David and Lucile Packard Foundation and from the Simons Foundation (SCOL #321183). J.H. acknowledges support from the National Science Foundation Graduate Research Fellowship under grant No. DGE-1144152. E.A.B. acknowledges funding through NSF grant AST-1514670 and NASA NNX16AB48G. C.W. acknowledges financial support from STFC (grant reference ST/R000549/1) and the University of Leeds.The volatile contents of protoplanetary disks both set the potential for planetary chemistry and provide valuable probes of defining disk system characteristics such as stellar mass, gas mass, ionization, and temperature structure. Current disk molecular inventories are fragmented, however, giving an incomplete picture: unbiased spectral line surveys are needed to assess the volatile content. We present here an overview of such a survey of the protoplanetary disks around the Herbig Ae star MWC 480 and the T Tauri star LkCa 15 in ALMA Band 7, spanning ∼36 GHz from 275 to 317 GHz and representing an order of magnitude increase in sensitivity over previous single-dish surveys. We detect 14 molecular species (including isotopologues), with five species (C34S, 13CS, H2CS, DNC, and C2D) detected for the first time in protoplanetary disks. Significant differences are observed in the molecular inventories of MWC 480 and LkCa 15, and we discuss how these results may be interpreted in light of the different physical conditions of these two disk systems.PostprintPeer reviewe
UV-driven Chemistry as a Signpost for Late-stage Planet Formation
The chemical reservoir within protoplanetary disks has a direct impact on
planetary compositions and the potential for life. A long-lived carbon-and
nitrogen-rich chemistry at cold temperatures (<=50K) is observed within cold
and evolved planet-forming disks. This is evidenced by bright emission from
small organic radicals in 1-10 Myr aged systems that would otherwise have
frozen out onto grains within 1 Myr. We explain how the chemistry of a
planet-forming disk evolves from a cosmic-ray/X-ray-dominated regime to an
ultraviolet-dominated chemical equilibrium. This, in turn, will bring about a
temporal transition in the chemical reservoir from which planets will accrete.
This photochemical dominated gas phase chemistry develops as dust evolves via
growth, settling and drift, and the small grain population is depleted from the
disk atmosphere. A higher gas-to-dust mass ratio allows for deeper penetration
of ultraviolet photons is coupled with a carbon-rich gas (C/O > 1) to form
carbon-bearing radicals and ions. This further results in gas phase formation
of organic molecules, which then would be accreted by any actively forming
planets present in the evolved disk.Comment: Accepted to Nature Astronomy, Published Dec 8th 202
Molecules with ALMA at Planet-forming Scales (MAPS). VI. Distribution of the small organics HCN, C2H, and H2CO
Funding: I.C. was supported by NASA through the NASA Hubble Fellowship grant HST-HF2-51405.001-A awarded by the Space Telescope Science Institute, which is operated by the Association of Universities for Research in Astronomy, Inc., for NASA, under contract NAS5-26555. C.W. acknowledges financial support from the University of Leeds, STFC, and UKRI (grant Nos. ST/R000549/1, ST/T000287/1, and MR/T040726/1). J.D.I. acknowledges support from the Science and Technology Facilities Council of the United Kingdom (STFC) under ST/T000287/1.Small organic molecules, such as C2H, HCN, and H2CO, are tracers of the C, N, and O budget in protoplanetary disks. We present high-angular-resolution (10-50 au) observations of C2H, HCN, and H2CO lines in five protoplanetary disks from the Molecules with ALMA at Planet-forming Scales (MAPS) ALMA Large Program. We derive column density and excitation temperature profiles for HCN and C2H, and find that the HCN emission arises in a temperate (20-30 K) layer in the disk, while C2H is present in relatively warmer (20-60 K) layers. In the case of HD 163296, we find a decrease in column density for HCN and C2H inside one of the dust gaps near ~83 au, where a planet has been proposed to be located. We derive H2CO column density profiles assuming temperatures between 20 and 50 K, and find slightly higher column densities in the colder disks around T Tauri stars than around Herbig Ae stars. The H2CO column densities rise near the location of the CO snowline and/or millimeter dust edge, suggesting an efficient release of H2CO ices in the outer disk. Finally, we find that the inner 50 au of these disks are rich in organic species, with abundances relative to water that are similar to cometary values. Comets could therefore deliver water and key organics to future planets in these disks, similar to what might have happened here on Earth. This paper is part of the MAPS special issue of the Astrophysical Journal Supplement.Publisher PDFPeer reviewe
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